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The genus Adenia Forssk., with ± 100 Old World spe-cies, occurs particularly in tropical and subtropical regionsof Africa, Madagascar, Southeast Asia, Malaysia and north-ern Australia (Feuillet & MacDougal 2007). A significantnumber of the African taxa are also associated with veryarid regions (e.g. Somalia and Namibia) (see Van Wyk &Smith 2001: 157, 158 on the Afro-arid cOITidorthat linksnorthern Namibia to the Horn of Africa). Ten species occurin southern Africa; most are dioecious, with monoecy orpolygamy rare (Archer 2000; Feuillet &MacDougal 2007).
Subsequent to publication of the typical subspeciesof Adenia fruticosa Burtt Davy, sect. Microblepharis(Wight & Am.) Engl. (Burtt Davy 1926), based only onmale material, Liebenberg (1939) described the femaleflowers. The revision of South African adenias by himaccommodated a broad concept for A. fruticosa, uphelduntil two further subspecies were distinguished by DeWilde (1971). For the delimitation of A.ji'uticosa BurttDavy subsp. simplicifolia De Wilde, herbarium mate-rial of both male and female flowers were available toits author. However, this was not the case for A. fruti-cosa Burtt Davy subsp. trifoliolata De Wilde, for whichfemale flowers and fruit were reportedly not seen. DeWilde (1971), nevertheless distinguished this KwaZulu-Natal endemic based on vegetative and male floral char-acters, and provided a key to this end, which was laterreproduced (De Wilde 1976) in slightly modified form.Information on fruits was additionally documented forthe Flora of southern Afi'ica account (De Wilde 1976),but not of female flowers, which were unavailable at thetime. We have been unable to trace the existence of anyhistorical vouchers representing female flowers.
During a field trip to Ulundi in Zululand in August2008, both male and female plants of this narrowly dis-tributed KwaZulu-Natal endemic were encountered infull and synchronous bloom; as is usual with woodydioecious taxa, female plants were far less prevalent thanmales in the subpopulation. A female voucher and furthermale herbarium specimens were gathered, enabling illus-
tration (Figure 21) and completion of the description forAdenia ji'uticosa subsp. trifoliolata, including an ampli-fied circumscription of the male element. Female floralcharacters (Table 3) confirm the distinctions recognizedby De Wilde (1971), for flowers of subsp. trifoliolataopen substantially wider than those of other subspecies,based in part on their somewhat longer sepals. Newlyacquired data have enabled the construction of a key forfemale plants of the three allopatric subspecies, all ofwhich occur within the FSA region (Figure 22). Adeniafruticosa subsp. simplicifolia occurs also in southernand eastern Zimbabwe and neighbouring Mozambique(De Wilde 1971,2002). The subspecific epithets of twoof the three taxa are misnomers: subsp. simplicifolia isnot always simple-leaved, and subsp. trifoliolata may be5-foliolate. Accordingly, leaf characters are not deemedparticularly useful when identifying material.
Field observations have revealed that flowering ofboth male and female plants of subsp. trifoliolata ex-tends, intermittently, from August to December, withfruiting occurring from September onwards.
la Leaves 3-5-foliolale; peliolule of leaflets 2-5(-7) mm;anthers ± 3.0 mm subsp.fi·uticosa
1b Leaves simple or 3(01' 4)-foliolate; leaflets sessile; anthers4.0-5.5 mm:
2a Hypanthium broadly cup-shaped, ± 5-saccate, corona hairs0.5-1.0 mm; disc-glands foliolate . . . . . subsp. simp/icifo/ia
2b Hypanthium cup-shaped, tapering, not saccate; coronahairs up to 0.5 mm, or partly absent; disc-glandsabsent subsp. trifoliolata
1a flowers opening to 12-14mm; stipe 1.5 mm ... subsp./rifo/iolara1b flowers opening to 6-7 mm; stipe up to I mm:2a Leaves 3-5-foliolate; petiolule of leaflets 2-5(-7) mm;
disc-glands absent; staminodes 3-4 mm ..... subsp.jl-u/icosa2b Leaves simple or 3-foliolate; leaflets sessile; disc-glands
0.2-0.5 mm; slaminodes 2-2.5 n1ln subsp. simp/icifolia
FIGURE 21.-Adeniafi·uticosa subsp. trifoliolata. A, habit; B, leaf; C, c3' flower (large form); D, c3' flower (small form); E, c3' half-flower diagram;F, petal: reduced lamina with awn; G, connective of c3' flower (arrowed); H, Cjl inflorescence with leaves; I, Cjl inflorescence structure; J, Cjlflower; K, Cjl half-flower diagram; L, ovule; M, mature fruit. Scale bars: A, M, 10 mm; B-E, H-J, 5 mm; F, G, 2 mm; K, I mm; L, 0.2 mm.Altist: Angela Beaumont.
CharacterStipe length (mm)Flower opening to (mm)Sepal length (mm)Petal length (mm)Staminode length (mm)Staminodes connate for (mm)Corona hair length (mm)Disc glands (mm)Pistil length (mm)Gynophore length (mm)Ovary (mm)Styles connate for (mm)Style arms (mm)Stigma diameter (mm)
sllbsp.ji·uticosa (De Wilde 1976)0.5-1.0
7
5-73-43-41-20.2absent6.5-8.0
2-33-4 x 3.0-3.5
0.70.5-0.7±I
The following descriptions and general notes for floralcharacters have been based on Ulundi material.
Stipe 1.5 mm long, shorter than in male flowers,articulation not clear; bract at base of stipe heart-shaped with tiny gland at base of lamina, I mm long;bracteoles 2, at apex of stipe, triangular, 0.5 mmlong. Hypanthium cup-shaped, base broadly cuneateto rounded, 2.5-3.0 mm from top of stipe to bases ofsepals, 2.8-3.8 mm wide. Sepals 5, arrangement quin-cuncial; innermost 2 sepals oblong, 7.5-8.0 x 2.5-3.0mm, midvein and immediate parallel, secondary, lat-eral veins somewhat thicker than rest of lamina, alto-gether forming a triangular thickened middle region,broadest at sepal base or sinus, narrowest at sepal apex,lamina without ornamentation except at region levelwith and immediately adjacent to apex of corona bear-ing some minute fimbriate processes; margins slightlyerose or sinuous, or entire basally, erose distally; apexacute, with pronounced beak-like, recurved extension;outermost 3 sepals oblong to slightly oblong-elliptic,6.0-8.5 x 2.5-3.0 mm, midvein thicker than lateraland immediate parallel, secondary, lateral veins, mid-vein and immediate secondary veins altogether lessthickened than in inner sepals, lamina without orna-mentation except at region level with and immedi-ately adjacent to apex of corona bearing some minutefimbriate processes, margins entire basally, slightlysinuous to entire distally. Disc-glands minute, 0.3 mmwide, at bases of sepals, level with base of fused partof gynophore-staminode structure. Petals 5, shorterthan sepals, inserted in sinuses of sepals, linear-Ianceo-late, 4.5-5.0 x 0.5-0.8 mm, 1-3-nerved; margins entirebasally, erose to slightly sinuous distally; base truncate;apex acuminate. Androecium of 5 staminodes, basesfused into a tube around gynophore, free parts of stami-nodes 2.25 mm long; bases broad, 2 mm wide; apexminute with tiny, incurved tip representing vestigialanther. Corona comprising ring of 5 vertical connec-tives, each one between base of petal and inter-starn i-nodal part of androecial column, connectives 0.5 mmlong, 0.5 mm wide, (when viewed from above), edgesfimbriate. Gynoecium: pistil 8 mm long from base offree gynophore to top of stigmatic arms; gynophore 3mm long from base of fused part with androecial col-
sllbsp. simplicifoLia (De Wilde 1976)0.5-0.7565-63-42.0-2.510.50.2-0.55.0-6.51.5-2
1.5-3 x 1.5-2.50.5-0.70.5-1.0± 1.5
subsp. trifoLioLata1.5
12-146.0-8.53-42.251.8
0.35
3.75 x 3I
0.5-1.0± 1.6
umn to base of ovary, free part of gynophore 1.5 mmlong; ovary 3.5 mm from base of gynophore to basesof styles, 3 mm diam.; placentation parietal, placentas3; ovules usually 6 per ovary, 2 per placenta, 0.5 mmlong, anatropous, with prominent ridge along lengthand beak-like apex, funiculus somewhat expanded;styles 3, connate for 0.5 mm, stylar anns 0.7-1.0 mmlong, each stylar arm split in 2; stigmas 3, vertical (i.e.parallel with long axis of flower), each connecting thetwo split stylar arms of each of 3 styles, 1.50 x 1.75mm, surface papillose, edges laciniate-papillose. Fruitsubglobose to broadly turbinate, 16.5 x IS mm, withpersistant remnants of perianth. Seeds (immature) withpitted testa and swollen funiculus next to base of seed.Figure 21H-M.
Hypanthium base to articulation, 1-3 mm long. Pedi-eel 2-7 mm long, 1 or 2 buds along pedicel, buds eachwith bract and bracteoles, bracts of buds minute, 1.0-1.8mm long, leaf-like; bracteoles of buds 2, I mm long,triangular margins irregular, articulation between baseof hypanthium and pedicel distinct; fused portion ofbases of sepal lobes cup-shaped in outline, base broadlycuneate to rounded, 2.5-3.5 mm deep (i.e. from pointof divergence of sepal lobes to apex of hypanthium),4.2-6.5 mm wide. Sepals quincuncial arrangement,inner 2 sepals oblong, 9.0-l2.8 x 3.0--4.8 mm, midveinsprominently thickened into a broad-based triangle, nar-rowing towards apex, margins entire or minutely wavybasally, minutely erose to unevenly serrate distally, apexacute to rounded, tip incurved, beak-like with fimbriae;outer 3 sepals oblong, 10-13 mm x 3.0--4.8 mm, mid-veins slightly thickened, margins entire to very slightlysinuous throughout, apex acute to rounded. Petals usu-ally 5, shorter than sepals, inserted in sinuses of sepallobes, development variable, linear-lanceolate, 5.0-8.8x 0.7-2.3 mm; margins slightly serrate to entire basally,erose to unevenly serrate distally; tips acuminate; laminasometimes partly reduced with awn-like extension ofmidvein, or lamina absent and petal represented by awn-like structure alone. Stamens 5, opposite sepal lobes; fil-aments broadly triangular, 2.5--4.5 x 0.5-1.0 mm, antherattachment sub-basal; anthers oblong, 3.0-5.0 x 0.5-1.8mm, bi-thecate, dehiscence introrse, pollen yellow.Corona connecting bases of petals to bases of filaments,
FIGURE 22.-Known distribution of Adeniaji'uticosa based on speci-mens at NH, NU and PRE and those cited by De Wilde (1971):subsp.fruticosa, .; subsp. trifoliolata, "'; subsp. simplicifolia,
••outermost (visible) tips of connectives of corona up to0.5 mm wide, minutely fimbriate, some fimbriae alsoarising from the inner surfaces of the sepals immediatelyadjacent to, and level with the tips of the corona arms;pistillode vestigial, up to 1 mm long. Figure 2IA-G.
Male flowers produced synchronously on single plantswere dimorphic in respect of overall flower size (Figure21C, D) and variable petal development as describedcurrently. De Wilde (1971) noted that considerable infra-specific variation in both the size of flowers, and theircomponents, is known for Adenia. Whereas Liebenberg(1939) related various abnormalities and variations inthe flowers of several South African genus members,he did not document male flower dimorphism in Adeniafruticosa.
Specimens examinedKWAZULU-NATAL.-283I (Nkandla): Ondini Historic Reserve,
Ulundi, male plant, 513 m, S 28° 18.765', E 31° 27.407', (-AD), 17-08-2008,1. van Vuuren 1 (NH); Ondini Historic Reserve, Ulundi, femaleplant, 515 m, S 28° 18.783', E 31° 27. 402', (-AD), 17-08-2008,1. vanVuuren 2 (NH).
James van Vuuren of Ulundi kindly brought the exist-ence of a subpopulation of Adenia ji-uticosa subsp. tri-foliolata to our attention. He subsequently collectedvoucher material of both sexes, and made observationson the phenology. The use in part of PRECIS data isgratefully acknowledged, kindly supported by MrsH. Snyman. The staff of the Mary Gunn Library at theNational Herbarium in Pretoria generously assisted withsourcing literature.
ARCHER, R.H. 2000. PassiAoraceae. In O.A. Leistner, Seed plants ofsouthern Africa: families and genera. Strelitzia 10: 434-436.National Botanical Institute, Pretoria.
BURTT DAVY, J. 1926. A manual oftheflowering plants andferns ofthe Transvaal with Swaziland, South Afi'ica, Pal1 I. Pteridophytato Bombacaceae: 36. Longmans, Green, London.
DE WILDE, WJ.J .0. 1971. A monograph of the genus Adenia Forsk.(PassiAoracaeae). Mededelingen Landbouwhogeschool Wage-ningen71,18: 1-281.
DE WILDE, WJ.J.O. 1976. PassiAoraceae. In J.H. Ross, Flora ofsouth-ern Afi'ica 22: 104-128. Botanical Research Institute, Pretoria.
DE WILDE, WJ.J.O. 2002. Passifloraceae. In U. Eggli, l!Ius/ratedhandbook on succulent plants: dicotyledons: 336-350. Springer-Verlag, Berlin.
FEUILLET, C. & MACDOUGAL, J.M. 2007. PassiAoraceae. In K.Kubitzki, Thefamilies and genera of vascular plants. IX Flow-ering plants-Eudicots: 270-281. Springer- Verlag, Berlin.
LIEBENBERG, L.C.C. 1939. A revision of the South African species ofAdenia. Bothalia 3: 513-570.
VAN WYK,A.E. & SMITH, G.F. 2001. Regions offloristic endemism insouthern Aji·ica. A review with emphasis on succulents. UmdausPress, Hatfield, Pretoria.
* Ethnobotany Unit, South African National Biodiversity Institute, P.O.Box 52099, 4007 Berea Road I School of Chemistry, University ofKwaZulu-Natal, 4041 Durban. Email: [email protected] (correspond-ing author).** School of Biological and Conservation Sciences, University ofKwaZulu-Natal, Private Bag XOI, 3209 Scottsville. Email: [email protected].*** Research and Scientific Services, South African National Biodi-versity Institute, Private Bag X I0],0001 Pretoria / Acocks Chair, Sch-weickerdt Herbarium, Department of Botany, University of Pretoria,0002 Pretoria. Email: [email protected]. received: 2009-06-05.
